Northeast Atlantic Late Quaternary planktic Foraminifera as primary productivity and water mass indicators

Transcription

1 Northeast Atlantic Late Quaternary planktic Foraminifera as primary productivity and water mass indicators Shirley A. van Kreveld Kreveld, S.A. van. Northeast Atlantic Late Quaternary planktic Foraminifera as primary productivity and water mass indicators. Scripta Geol., 113: 23-91, 18 figs., 12 pls, Leiden, December S.A. van Kreveld, Center for Marine Earth Sciences, Institute of Earth Sciences, Free University, De Boelelaan 1085, 1081 HV Amsterdam, The Netherlands. Keywords: Planktic Foraminifera, palaeo primary productivity, water mass, Northeast Atlantic. Primary productivity and water mass reconstructions based on planktic Foraminifera reveal distinct interglacial/glacial variations for the past 208 ka in a mid-latitude Northeast Atlantic piston core. Average total planktic foraminiferal absolute frequencies and accumulation rates, which are interpreted to reflect primary productivity, are higher in interglacial than in glacial sediments. Low total planktic foraminiferal absolute frequencies and accumulation rates in 'Heinrich layers' are likewise interpreted to show low production of Foraminifera due to low surface ocean fertility. 'Heinrich layers' are enriched in ice-rafted debris, recording periods of massive iceberg production and rapid melting in the Northeast Atlantic. The dominance of Neogloboquadrina pachyderma (sinistral) in these layers reflects an extension of cold low salinity polar waters. The fresh water along with turbidity caused by melting icebergs may account for the low productivity during these events. In contrast, the dominance of Globigerina bulloides, Neogloboquadrinaincompta,Globorotalia scitula, Globigerinita glutinata, and Globorotalia inflata group in interglacial sediments is interpreted to reflect conditions comparable with the present day North Atlantic Current (NAC) waters in the area. In the modern ocean, the Gulf Stream and its extension, the NAC, are driven by seasonally strong westerly winds which induce mixing, supplying nutrients from deep to surface waters. Enough food and sufficient light combine to provide for pulses of algal blooms which support large populations of Foraminifera. The important planktic foraminiferal test contributors in glacial sediments are not only Turborotalita quinqueloba, a subpolar species, but also polar and NAC indicator species. Primary productivity is inferred to be extremely seasonal, with low productivity during winter when there was little sunlight and partial ice cover. Planktic Foraminifera are good primary productivity indicators in this carbonate-dominated, openocean Northeast Atlantic site. In contrast, organic carbon, which is directly linked to primary productivity and is extensively used as a proxy in upwelling areas, is unsuitable in this site because of the very low organic carbon content of the sediments and the contamination by terrigenous organic matter. Contents Introduction 24 Hydrography 25 Material and methods 26 Results 28 Stratigraphy 28 Age model 30 >125 urn planktic foraminiferal frequencies and accumulation rates 33 Organic carbon 35 Cluster and correspondence analyses 35

2 24 van Kreveld. Planktic Ibraminifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) Diversity indices 47 Discussion and interpretation 47 Palaeo primary productivity 47 Planktic Foraminifera 47 Organic carbon 51 Water masses 52 Productivity and water masses 54 Conclusions 54 Acknowledgements 56 References 56 Appendix I 63 Introduction The observed variability in atmospheric C0 2 concentrations during the Late Quaternary (Delmas et al., 1980; Barnola et al., 1987; Neftel et al., 1988) is attributed, at least in part, to changes in surface water productivity and ocean circulation (Broecker, 1992). Primary producers, such as coccolithophorids, can act as a 'biological pump' (Berger et al., 1987) by drawing down both surface water and atmospheric C0 2. Since biological productivity and ocean circulation are important in redistributing carbon in the ocean, palaeo primary productivity and surface water mass changes are reconstructed during the last glacial cycle in a mid-latitude open-ocean site. Although productivity is lower than in coastal upwelling areas, the large areal coverage of the open-ocean makes it an important factor in the global carbon budget (Milliman, 1993). These reconstructions are valuable in constraining global carbon cycle models. This paper aims to reconstruct glacial to interglacial palaeo primary productivity and surface water mass variations in mid-latitude Northeast Atlantic using planktic foraminiferal accumulation rates and relative frequencies, respectively. For this purpose, I studied piston core T88-9P because it has a 208 ka sediment record and was unaffected by dissolution as it was recovered way above the lysocline. Moreover, this core is ideal for this study because it was recovered in mid-ocean where extensive coccolithophorid blooms presently occur (Brown & Yoder, 1994a), outside the influence of turbidity currents and at an optimum position for receiving debris discharged during iceberg melting. Planktic Foraminifera can be used to reconstruct palaeo primary productivity variations from deep-sea cores because their fluxes are strongly correlated with surface ocean productivity (Thunell & Reynolds, 1984). Additionally, foraminiferal assemblages characterise distinct biogeographic zones which are intimately linked to particular water masses (Boltovskoy, 1959, 1962; Bé & Tolderlund, 1971; Vincent & Berger, 1981; Ottens, 1991), and as the tests of foraminiferal species are preserved in the sediments, they can be used to reconstruct past water mass changes. These were reconstructed by grouping the relative frequencies of the planktic foraminiferal species using cluster and correspondence analyses and then comparing these groups to present day plankton distribution and their associated water masses. Diversity indices were also calculated to aid in water mass reconstructions.

3 van Kreveld Planktic R>raminifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 25 Aside from planktic Foraminifera, the organic carbon content of the sediments, which is primarily derived from biological production in surface waters, has also been extensively used as a primary productivity indicator (Müller & Suess, 1979; Betzer et al., 1984; Stein & Stax, 1991; Thunell et al., 1992; Sarnthein et al., 1992). In this paper, the usefulness of organic carbon as a fertility proxy in an open-ocean, mid-latitude Northeast Atlantic site is assessed. Planktic foraminiferal assemblages preserved in deep-sea sediments have been extensively used to reconstruct (ecologie) water mass movements (CLIMAP, 1976, 1984). They were also used as a primary productivity proxy. For example, high frequency of Globigerina bulloides was utilised as an index for high productivity (Prell, 1984a) whilst the ratio of Neogloboquadrina dutertrei to Pulleniatina obliquiloculata was used as a qualitative fertility indicator (Berger & Killingley, 1977). Mix (1989) and Sarnthein et al. (1992) extended the one species based index to all the planktic Foraminifera by using transfer functions to arrive at quantitative estimates of primary production. The above proxies only use relative frequencies of planktic Foraminifera. This study is among the few providing data on planktic foraminiferal accumulations rates which can directly be compared with sediment trap flux data. Other organisms, including benthic Foraminifera (Thomas et al., 1995; Herguera & Berger, 1991; Herguera, 1992, 1994; Berger et al., 1994), coccolithophorids (Pujos, 1992; Giraudeau & Rogers, 1994), diatoms (Sancetta, 1992; Schräder, 1992; Abrantes et al., 1994; Takahashi, 1994), and dinoflagellates (Lourens et al., 1992; Dale & Fjellsa, 1994; Versteegh, 1995), also serve as primary productivity proxies. Aside from microfossils, chemical tracers, such as carbonate (Brummer & van Eijden, 1992; van Kreveld et al., 1996), opal (Mortlock et al., 1991), barium (Dymond et al., 1992), and oxygen and carbon isotopes in planktic Foraminifera (Steens et al., 1992) have been also utilised as indicators of past surface water productivity. Hydrography The present day circulation of the North Atlantic is characterised by the southward flow of the cold East Greenland Current and the Labrador Current and the northeastern flow of the Gulf Stream (Fig. 1). The cold, low saline Labrador Current joins the warm saline extension of the Gulf Stream off Newfoundland to form the North Atlantic Current (NAC) (Dietrich et al., 1975; Krauss, 1986). The northern NAC component crosses the Mid-Atlantic Ridge at 52 N into the Northeast Atlantic (Krauss, 1986) (Fig. 1). The southern branch of the NAC heads towards the Azores and continues to Madeira (Mann, 1967; Käse & Siedler, 1984; Siedler et al., 1985). A transitional zone separates the anticyclonic (clockwise) gyre of subtropical waters to the south from the cyclonic (counterclockwise) gyre in the north (Krauss, 1986). The surface water circulation is associated with polar, subpolar, NAC, transitional, and subtropical water masses (Fig. 1) (Krauss, 1986). The southern limit of the East Greenland Current approximately marks the present day polar front. The northern limit of the NAC approximates the subpolar front while its southern limit is associated with a transition zone, separating the subtropical waters to the south from the northern water mass (Krauss, 1986). In contrast to modern times, oceanic surface water reconstructions for the last glacial maximum (Ruddiman & Mclntyre, 1976; CLIMAP, 1981), show that ice-bear-

4 26 van Kreveld. Planktic Foraminifera as primary productivity and water mass indicators,scriptageol, ) Fig. 1. Océanographie map of the Northeast Atlantic Ocean depicting present day surface currents (Krauss, 1986) and their associated water masses. Filled circle shows the position of piston core T88-9P (48 23'2" N, 25 05'6" W; water depth = 3193 m; core length = 7.90 m). NAC stands for North Atlantic Current. ing polar water covered the North Atlantic above latitude 45 N in a cold counterclockwise gyre (Fig. 2). A west-east oriented subpolar front developed at this latitude parallel to the eastern path of the Gulf Stream (equivalent to NAC) which was compressed to a narrow band between the clockwise subtropical gyre and the counterclockwise polar gyre (Fig. 2). Material and methods Piston core T88-9P ( N, W; recovery 790 cm) (Fig. 1) provides a, M, M good planktic foraminiferal record for the last 208 ka since it is well- preserved, continuous, and recovered well above the lysocline at 3193 m water depth. This core has undergone minimal carbonate dissolution as shown by the presence of aragonitic pteropod shells as well as fragile and juvenile planktic Foraminifera specimens with spines occasionally preserved, even during isotopic stage 2 when carbonate dissolu-

5 van Kreveld. Planktic Foraminifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 27 Fig. 2. Palaeoceanographic map of the Northeast Atlantic ocean showing inferred surface currents and water masses during the last glacial maximum (redrawn from Ruddiman & Mclntyre, 1976 and CLI MAP, 1981). Bold lines show the ice extent. tion was high (Crowley, 1981). The core was collected in April, 1988 during the 'Actuomicropalaeontology Palaeoceanography North Atlantic Project' (APNAP- II) R/V Tyro cruise. It was sampled every 5 cm in the upper 50 cm and then at about 10 cm intervals. These 10 cm, 3 1 cm thick samples, totalling 83, were sieved through 125 urn mesh screens and the fractions were weighed. The >125 urn fraction was repeatedly split to obtain an aliquot of c. 300 specimens of planktic Foraminifera. There were 16 species found and the most common species are illustrated in plates One sample (238 cm) did not contain any planktic Foraminifera. The weight % carbonate of the bulk sediment was analysed with a ±2% precision by a modified Scheibler-type gasometric technique (Bruin, 1937). Gray reflectance profiles were obtained by digitally scanning and measuring the relative reflectivity of the coloured core photograph (courtesy of L. Labeyrie and E. Cortijo, Centre des Faibles Radioactivités, Laboratoire mixte CNRS-CEA, Gif sur Yvette Cedex, France).

6 28 van Kreveld. Planktic Foraminifera as primary productivity and water mass indicators,scriptageol., ) Dry bulk density was determined by weighing 5 cm of sediment after drying at 3 50 C. Three samples weighing c. 15 mg of >250 urn sized mixed planktic Foraminifera were dated by radiocarbon measurements using accelerator mass spectrometry (AMS). Forty-six bulk samples were analysed in duplicate for organic carbon content using a Carlo Erba NA-1500 elemental analyser according to the method described by Nieuwehuize et al. (in press). These samples were oven-dried at 60 C, then homogenised by thorough grounding in an agate mortar mill to reduce variability between duplicates. The organic carbon content was measured after removal of the inorganic carbon by acidification using 25% HCl in silver sample cups. Duplicate measurements were within ± 0.06%. The accumulation rates of planktic Foraminifera were calculated using the following formulas: (1) pftar = S*DBD*pf/g (2) pfsar = S*DBD*pf species/g whereby: pf/g=planktic Foraminifera per gram dry weight (no./g); pftar=total planktic foraminiferal accumulation rate (no./cm ka); 2 pfsar=planktic foraminiferal species accumulation rate (no./cm ka); 2 S=sedimentation rate (cm/ka); DBD=dry bulk density; dry weight per wet volume (g/cm ); 3 Cluster and correspondence analyses were used on the planktic Foraminifera relative frequency data matrix to determine systematic faunal changes through time. Both samples and species relative frequency were clustered separately using a weighted pair group cluster method on a correlation matrix (Davis, 1986). Correspondence analysis (Hill, 1974; Davis, 1986; Bénzecri, 1992) is one of several ordination methods which reduces the dimensionality of large data matrices. It extracts eigenvectors from a chi-square weighted distance matrix calculated from the relative frequencies of the planktic foraminiferal species and the samples which are taken from different depth intervals in the core. Correspondence analysis has the advantage that samples and species scores are scaled so that both can be plotted in one graph, thus facilitating interpretation. Diversity indices were calculated to document diversity trends through time. Simple diversity gives the number of species in a sample, the Shannon diversity index expressed as (H'=- p*\npù takes into account the relative proportion of each species (pi) in a sample (Shannon, 1949), and equitability (E'=e '/s in which s is the H number of species in a sample) measures the evenness of the species distribution in a sample (Buzas & Gibson, 1969). When one species dominates the foraminiferal assemblage, equitability approaches zero, while it is one when all species are present in equal proportions. Results Stratigraphy The interglacial sediments of piston core T88-9P are coccolith and foraminiferal oozes whilst the glacial sediments are mostly calcareous sand, silt and clay (Fig. 3A).

7 van Kreveld Planktic R)raminifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 29 Fig. 3. Core T88-9P (A) lithologie column based on visual core description, (B) weight % carbonate content of the bulk sample and (C) gray reflectance profile plotted against depth. Light-coloured sediments have higher gray reflectances than dark-coloured ones, as shown by the higher numbers. Rectangles hi- hi3 are intervals interpreted as Heinrich layers.

8 30 van Kreveld. Planktic Foraminifera as primary productivity and water mass indicators,scriptageol, ) Thus, the former generally have a higher weight % bulk carbonate (up to 87%) while it can be as low as 7% in the latter (Fig. 3B). As a result of the varying carbonate content, the interglacial layers are cream to brown coloured and usually have higher gray reflectances than glacial layers which are olive gray to gray (Fig. 3C). 'Heinrich layers' are considered to record massive influxes of icebergs into the North Atlantic (Heinrich, 1988; Broecker et al., 1992; Bond et al., 1992; Grousset et al., 1993). Van Kreveld et al. (1996) identified thirteen intervals in the core which were interpreted as Heinrich layers based mainly on the high ice-rafted debris content and the presence of detrital carbonate. These layers also have very low planktic foraminiferal accumulation rates (>125 urn) with N. pachyderma sinistral (s) dominating the foraminiferal assemblage (van Kreveld et al, 1996). Most of these layers contain dropstones and have high dry bulk densities. The Heinrich layers in core T88-9P were successively numbered from top to bottom and designated with a small letter h. The upper six correspond to those identified by Heinrich (1988). However, the lower seven identified in isotopic stages 6-7 are different from those of Heinrich (1988) which are in stage 5. Van Kreveld et al. (1996) discuss the differences in the timing of the Heinrich layers identified in nearby cores. Heinrich layers hi, h2, h4 and h5 have higher gray reflectances than h3 (Fig. 3C), due to their higher detrital carbonate content. The detrital carbonate originates from limestone and dolomite bedrocks in eastern Canada (Andrews & Tedesco, 1992; Grousset et al., 1993). Age model The ages are based on three radiocarbon dates, on two ash layers and on the correlation of the oxygen isotope curves of Globigerina bulloides and Globorotalia inflata with the chronostratigraphy of Martinson et al. (1987) (Fig. 4A-C) (van Kreveld et al., 1996). The radiocarbon age of the surface sample was calibrated to a calendar date using the program Calib 3.0 (Stuiver & Reimer, 1993) which is based on the marine bidecadal data set of Stuiver & Braziunas (1993) while those of samples 128 cm ( ± 300 years BP) and 148 cm ( ± 300 years BP) were calibrated using Th-U ages obtained by mass spectrometry on corals (Bard et al., 1990,1993) after subtracting 400 C years for reservoir correction (Bard, 1988). 1 4 The chronology of the core was supplemented by the carbonate content percentage, N. pachyderma (s) frequency, coccolith abundance and gray scale records to identify isotopic stage boundaries 5/4 and 4/3.1 also used the strong gradient in the carbon isotope curves of both Globigerina bulloides and Globorotalia inflata to identify the 5/4 boundary (Fig. 5A). The depth to age conversion was modelled by linearly interpolating between the age control points by assuming uniform sedimentation rates to obtain the date for each sample (Fig. 5B). The sedimentation rates, which are generally higher for glacial than for interglacial sediments, vary from 2.2 cm/ka to 7.5 cm/ka. I assume that the Heinrich layers have the same sedimentation rate as the surrounding glacial sediments, therefore the planktic foraminiferal accumulation rate in Heinrich layers is a minimum estimate while it is a maximum estimate for glacial sediments. This

9 van Kreveld Planktic Foraminifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 31 Fig. 4. (A) Globigerina bulloides, Globorotalia inflata and Neogloboquadrina pachyderma (s) oxygen isotope ratios. (B) Control points for depth to age transformation are mainly based on calendar-calibrated radiocarbon ages, ash layers and isotope stratigraphy. (C) Orbitally-based chronostratigraphy of Martinson et al. (1987), with the events and boundaries used for the age model. Stippled lines mark the isotopic stage boundaries.

10 32 van Kreveld. Planktic Foraminifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) Fig. 5. (A) Carbon isotope ratios of Globigerina bulloides and Globorotalia inflata versus depth. Depth to age plot showing the sedimentation rate (cm/ka) between age control points.

11 van Kreveld. Planktic Foraminifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 33 assumption is supported by more detailed dating in nearby cores (Bond et al., 1992; Francois & Bacon, 1994) of the upper three Heinrich layers, which show similar or only slightly higher sedimentation rates. The non-heinrich layers of isotopic stages 7, 5 and 1 will subsequently be referred to as interglacial, and that of isotopic stages 6 and 4-2 as glacial. >125 urn planktic foraminiferal frequencies and accumulation rates The total planktic foraminiferal absolute frequencies (number of planktic foraminiferal specimens per gram bulk sediment) are generally higher in interglacial than in glacial sediments, averaging and specimens/g, respectively (Table 1; Fig. 6A). An interval in h4 was even devoid of Foraminifera. The highest absolute frequency is found in isotopic stage 5. The total planktic foraminiferal accumulation rates show a similar pattern, which is higher in interglacial than glacial sediments, with a mean of and specimens/cm2ka, respectively (Table 1; Figs. 6B and 7B). The planktic foraminiferal assemblage in piston core T88-9P sediments is dominated by seven species, viz. N. pachyderma (s), Turborotalita quinqueloba, Globigerina bulloides, Neogloboquadrina incompta, Globorotalia scitula, Globigerinita glutinata, and Globorotalia inflata (Figs. 6C and 8). Interglacial deposits have a low average relative frequency of Neogloboquadrina pachyderma (s) and generally high to moderate percentages of the other species. Glacial sediments have high relative frequencies of N. pachyderma (s) and T. quinqueloba, and moderate to low percentages of the other species while Heinrich layers are dominated by N. pachyderma (s). Nine other species which do not exceed 4% of the total, include Globorotalia hirsuta, Globigerinella aequilateralis, Globigerinoides ruber, Orbulina universa, Globorotalia truncatulinoides, Globigerina rubescens, Globigerinoides tenellus, Turborotalita humilis and Globigerinita uvula (Figs. 6C and 8). These species are generally found in interglacial sediments and are rarely present in glacial deposits and Heinrich layers. Neogloboquadrina pachyderma (s) averages 8.7% of the total planktic foraminiferal assemblage in interglacial, 44.6% in glacial, and 57.7% in Heinrich layers, corresponding to mean absolute frequencies of 3300, 9000, and 4400 specimens/g, respectively (Table 1; Figs. 6A and C). Although N. pachyderma (s) dominates in Heinrich layers, its absolute frequency is generally lower than in glacial sediments. It does not exceed 2500 specimens/g in the upper five Heinrich layers (hl-h5) though it may comprise up to 95% of the total foraminiferal fauna (compare Figs. 8 and 9). The absolute frequency of N. pachyderma (s) is generally high immediately before and after a Heinrich layer. The accumulation rate of N. pachyderma (s) is at least three times lower in interglacial than in glacial sediments (Fig. 10). Although the average absolute frequency and accumulation rate of T. quinqueloba are higher in interglacial than in glacial sediments, its average relative frequency is slightly higher in the latter (Table 1; compare Figs. 6A and C). The average frequencies of this species are low in Heinrich layers (Figs. 8 and 9). The average relative frequencies of G. bulloides are high (16.7%), moderate (13.4%), and low (7.3%) in interglacial, glacial and Heinrich layers corresponding to mean absolute frequencies of 5900, 2900 and 900 specimens/g, respectively (Table 1;

12 34 van Kreveld. Planktic Foraminifera as primary productivity nd water mass indicators, Scripta GeoL, ) Fig. 6. (A) Absolute frequency (no.*1000/gram), (B) accumulation rate (no.*1000 /cm 2 ka) and (C) relative frequency (%) averages of the >125 urn planktic foraminifera in interglacial, glacial and Heinrich layers. Figs. 6A and C). This species does not exceed 100 specimens/g in Heinrich layers hlh5, even though it may comprise up to 20% of the planktic foraminiferal assemblage (Figs. 8 and 9). G. bulloides has a higher interglacial than glacial accumulation rate (Fig. 10). Of all the planktic Foraminifera, N. incompta has the highest average frequencies and accumulation rate in interglacial sediments. Its average accumulation rate is more than two times higher in interglacial than in glacial layers (Table 1; Figs. 6B and 10). Among the seven dominant species, G. scitula has the lowest average frequencies of only 5% and 1800 specimens/g in interglacial layers (Table 1, Figs. 6A and C). It is less frequent in glacial and Heinrich layers. The average accumulation rate of G. scitula is at least three times higher in interglacial than in glacial sediments (Figs. 6B and 10). Both average relative and absolute frequencies of G. glutinata are moderate, low and very low in interglacial, glacial and Heinrich deposits, respectively (Table 1; Figs. 6A and C). The average accumulation rate of this species is about three times higher in interglacial than in glacial sediments (Figs. 6B and 10).

13 van Kreveld. Planktic Forarnirdfera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 35 Globorotalia inflata shows decreasing average relative and absolute frequencies from interglacial, glacial to Heinrich layers of 9.7%, 2.9%, 1.8% and 3400, 600, 200 specimens/g, respectively (Table 1; Figs. 6A and C). The average accumulation rate of this species is about a factor four higher in interglacial than in glacial sediments. Organic carbon The total organic carbon content of the bulk sediment is very low, varying only from 0.06 to 0.22 % (Fig. 11). It is generally higher during isotopic stages 4 to 2 than in the other stages. Across Terminations I and II, and in most of the Heinrich layers the total organic carbon is low, except for Heinrich layers 3 and 6 which show peak maxima. Cluster and correspondence analyses Cluster analysis of the relative frequency data matrix shows five species clusters, I: N. pachyderma (s); II: T. quinqueloba; III: G. bulloides, N. incompta, G. scitula, G. glutinata, and G. inflata;yv: G. ruber, G. aequilateralis, and G. hirsuta; V: G. truncatulinoides, G. rubescens, G. tenellus, and O. universa (Fig. 12). Species with frequencies lower than 0.2% and the sample at 238 cm, which did not contain planktic Foraminifera, were excluded from all the analyses.

14 36 van Kreveld Plankticforaminiferaas primary productivity and water mass indicators, Scripta GeoL, ) Sample clustering also showed five groups (Figs. 13 and 14A-B). Cluster A consists mostly of Heinrich and isotopic stages 2 and 6, with some stage 3 samples. Cluster B mostly combines samples from isotopic stages 3 and 6, with some from 4 and Heinrich layers. Clusters C and E generally consist of samples from isotopic stages 5 and 7 while cluster D is dominantly made up of isotopic stage 1 samples. These five sample clusters, represent 34,11,19, 8, and 10 of the total 82 samples corresponding to 42,13, 23,10, and 12%, respectively. To relate species to sample clustering, the sum of the relative frequency of the species clusters in a sample (Fig. 14A) and the sample cluster designation (Fig. 14B) were plotted against time. In glacial intervals, cluster A is generally succeeded by cluster B and vice versa while during interglacial intervals, cluster C may either be succeeded by clusters D or E. Across Termination II, there is a drastic change from cluster I to V. Figure 14A also shows that the sum of the relative frequency of the species clusters IV and V in a sample is only <10%. Therefore, clusters IV and V are of less importance and the interpretations are concentrated on clusters I to III. The correspondence analysis more or less confirms the results of the cluster analysis though it is more difficult to interpret. The first correspondence analysis axis which is mainly controlled by N. pachyderma (s), explains 69% of the total variation (Figs. 15A and 16A). All other axes have only minor contributions (Figs. 15 B-C and 16A-B) although they distinguish groups that are similar to the clusters.

15 van Kreveld. Planktic Foraminifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 37 Table 1. Average relative frequency in %, absolute frequency in no./g and accumulation rate in no./cm 2 ka of the dominant planktic foramiferal species in interglacial, glacial and Heinrich layers. Average frequencies and accumulation rates of the dominant >125 urn planktic foraminiferal species. Average relative frequency (%) interglacial glacial Heinrich layers N. pachyderma (s) T. quinqueloba G. bulloides N. incompta G. scitula G. glutinata G. inflata Average absolute frequency (nc»-/g) interglacial glacial Heinrich layers Total planktic foraminifera N. pachyderma (s) T. quinqueloba G. bulloides N. incompta G. scitula G. glutinata G. inflata Average accumulation rate (no./cm ka) 2 interglacial glacial Heinrich layers Total planktic foraminifera N. pachyderma (s) T. quinqueloba G. bulloides N. incompta G. scitula G. glutinata G. inflata Figures 16 A-B show the plots of the first three main factors that control the data set. The coordinates of the individual species or samples are indicative of their contribution to the variation of the particular axis. Correspondence axes 1 vs 2 and axes 1 vs 3 show five distinct groups corresponding to the species cluster (compare Figs. 16 A-B and 12). Fig. 16B and 15C show high positive and negative loadings on axis 3 of the species belonging to clusters V and IV, respectively (compare with Fig. 12). However, because no samples fall within these species range, IV and V are only of minor importance.

16 38 van Kreveld Planktic Foraminifera as primary productivity and water mass indicators, ScriptaGeoL, ) Fig. 7. (A) >125 urn total planktic foraminifera per gram bulk sediment and (B) accumulation rate in no.*1000/cm 2 ka plotted against age.

17 van Kreveld Planktic Foraminifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 39 Fig. 8. >125 urn planktic foraminiferal species relative frequency in %, with the frequencies adding up to 100%. The water mass designation is based on clusters I-V of Fig. 12) with I-polar, Il-subpolar, Ill- North Atlantic Current, IV- transitional, and V-subtropical. 1-7 are isotopic stages.

18 40 van Kreveld. Planktic Foraminifera as primary productivity and water mass indicators,scriptageol, ) Fig. 9. >125 urn planktic foraminiferal species absolute frequency in no.*1000/g. 1-7 are isotopic stages.

19 van Kreveld Planktic R)raminifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 41 Fig. 10. >125 urn planktic foraminiferal species accumulation rate in no.*1000/cm 2 ka. 1-7 are isotopic stages.

20 42 van Kreveld. Planktic Foraminifera as primary productivity and water mass indicators,scriptageol, ) Fig. 11. Average weight percent organic carbon of the bulk sediment. Bars show duplicate measurements.

21 van Kreveld. Planktic Forandnifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 43 Fig. 12. Dendogram of the weighted pair group cluster analysis of the planktic foraminiferal species based on the correlation matrix of relative frequencies, excluding species lower than 0.2%. The five major clusters are interpreted to reflect water masses.

22 44 van Kreveld. Planktic Foraminifera as primary productivity and water mass indicators,scriptageol, ) Fig. 13. Dendogram of the weighted pair group cluster analysis of the counted planktic foraminiferal samples from piston core T88-9P based on the correlation matrix of relative frequencies, excluding sample number 238, which did not contain planktic foraminifera. Sample numbers give depth below the mudline in cm. Asterisks (*) mark Heinrich intervals.

23 van Kreveld Planktic Foraminifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 45 Fig. 14. (A) Sum of the relative frequency of the species clusters in a sample and (B) the sample cluster designation plotted against time.

24 46 van Kreveld Planktic Foraminifera asprimaryproductivityandwatermassindicators,scriptageol, )

25 van Kreveld Planktic Foraminifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 47 Fig. 15. Loadings of the different species on the first (A), second (B) and third (C) correspondence axes. Diversity indices In the past 208 ka, diversity was generally high except in Heinrich layers when simple diversity, Shannon and equitability indices have low values (Fig. 17). Both simple and Shannon diversity indices are generally slightly higher in interglacial than in glacial sediments. This trend is not reflected in the equitability. Figure 18 shows the inverse relationship between % N. pachyderma (s) and the Shannon diversity index. It also shows that the Shannon index is low to very low for species cluster I, moderate for cluster II and high for clusters III-V. Discussion and interpretation Palaeo Primary Productivity Planktic Foraminifera Although the sedimentary record of planktic foraminiferal assemblages represents a composite mixture of different seasons over many years, it is still possible to link it to general primary productivity patterns in the surface ocean. Food supply is crucial to foraminiferal abundance, therefore, variations in frequencies and accumu-

26 48 van Kreveld. Planktic Foraminiferaasprimaryproductivityandwatermassindicators,ScriptaGeoL, ) Fig. 16. Plot of samples and species on the (A) first vs second and (B) first vs third correspondence analysis axes. The first three axes explain 69,10 and 6% of the total variation in the data set. Samples belonging to the same cluster (also see Fig. 12) are given an identical symbol.

27 van Kreveld. Planktic Foraminirera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 49 Fig. 17. Diversity indices (A) simple, (B) Shannon and (C) equitability of planktic foraminifera, after (Buzas & Gibson, 1969). Diversity is generally very low in Heinrich layers. One sample at about 43 ka is devoid of foraminifera, thus the diversity indices are indeterminate.

28 50 van Kreveld. Planktic Foraminifera as primary productivity and water mass indicators,scriptageol, ) lation rates of planktic Foraminifera can be interpreted to reflect variations in primary productivity. This reasoning is supported by experimental data which showed increased food availability to enhance foraminiferal population growth (Bradshaw, 1955; Lipps & Valentine, 1970; Anderson et al., 1979; Bé et al., 1981; Caron et al, 1981, 1987). Additionally, planktic Foraminifera are plentiful in modern surface oceans where nutrients and biomass concentrations are high (compare Bé & Tolderlund, 1971 with Koblents-Mishke et al., 1970 and Berger et al., 1987 productivity maps; compare Bradshaw, 1959 with the productivity maps of Reid, 1962). The same phenomenon is observed in upwelling areas of the northern Indian Ocean (Kleijne et al., 1989). Sediment trap data also show that the fluxes of planktic Foraminifera are often strongly correlated with surface ocean productivity (Thunell et al., 1983; Thunell & Reynolds, 1984; Reynolds & Thunell, 1985; Thunell & Honjo, 1987). In this study, the generally higher total planktic foraminiferal absolute frequencies and accumulation rates during interglacial than during glacial periods are interpreted to reflect higher production of planktic Foraminifera in the surface waters due to increased primary productivity. Indeed, the estimate for the top sediment ( specimens of planktic Foraminifera >125 um/cm ka) compares well with flux data 2 for one year ( specimens of planktic Foraminifera >150 um/cm ka) from a sediment trap near the studied site (Wolfteich, 1994). These two estimates are compar 2 able despite the difference because the number of planktic Foraminifera in the urn fraction constitutes on average one third of the total planktic Foraminifera larger than 125 urn (see Peeters et al., in prep.). In the trap, the fluxes of planktic Foraminifera exhibit sharp increases during the spring bloom period with G. bulloides being the most dominant planktic foraminiferal species (Wolfteich, 1994). The coccolith carbonate accumulation rate is higher during interglacial than during glacial periods, which is similar to the pattern shown by the planktic Foraminifera (van Kreveld et al., 1996). These coccoliths are formed by coccolithophorids, which abound during phytoplankton blooms presently occurring in surface waters overlying the site in response to seasonal development and decay of the thermocline, and local upwelling within cyclonic eddies generated from meandering current systems of the North Atlantic Current (Wolfteich, 1994). The generally low total planktic foraminiferal absolute frequencies and low accumulation rates in Heinrich layers are interpreted to reflect depressed planktic foraminiferal productivity as a consequence of limited food supply in the photic zone. This interpretation is supported by the very low coccolith carbonate accumulation rate in the Heinrich layers, particularly in hi to h5 which are almost devoid of coccoliths (van Kreveld et al., 1996). During these events, the iceberg-laden waters probably blocked sunlight, slowing down algal and hence planktic foraminiferal production. Additionally, the lowered salinity due to the meltwater influx (Bond et al., 1992) and the cooling (Maslin et al, 1995; Madureira et al., in press) may have been unfavourable for the growth of coccolithophorids and Foraminifera. Thomas et al. (1995) also showed Heinrich events as periods of low primary productivity based on the very low benthic foraminiferal accumulation rates, and absolute and relative frequencies of the phytodetritus feeding species of benthic Foraminifera. Broecker et al. (1992) and Bond et al. (1992) also interpret Heinrich events to be periods of low primary productivity.

29 van Kreveld. Planktic Foraminifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 51 Aside from depressed primary productivity, they also give ice-rafted debris dilution as a possible alternative explanation for the lowered abundance of planktic Foraminifera in Heinrich layers. If so, then the sedimentation rate should at least be an order of magnitude higher than my estimates of between 5.7 to 7.5 cm/ka for the intense Heinrich events 1-3. However, detailed Th/U (Francois & Bacon, 1994) and radiocarbon dating (Bond et al., 1992) of cores recovered in the vicinity give similar values or just about 2 times higher, therefore dilution is unlikely to be the sole cause for planktic foraminiferal sparseness in these layers and decreased surface water productivity must have played a role. Moreover, the lateral extent of the area low in planktic Foraminifera exceeds that of the path of the Canada-derived debris (Broecker et al., 1992; Bond et al., 1992), supporting this contention. Higher interglacial than glacial primary productivity was also observed in the mid- to high-latitude North Atlantic using benthic Foraminifera (Thomas et al., 1995), carbonate mass accumulation rates (van Kreveld et al, 1996), and organic carbon content (Stein & Stax, 1991; van Weering & de Rijk, 1991) proxies. At high latitudes in the Southern Oceans productivity was also higher during interglacials (Berger & Wefer, 1991; Charles et al., 1991; Mortlock et al., 1991; Kumar et al., 1993; Francois et al., 1993; Shemesh et al., 1993). To the contrary, primary productivity was higher during glacial than interglacial periods in low latitude regions and upwelling areas of the Atlantic (Müller et al., 1983; Pokras, 1987; Sarnthein et al., 1992) and the Pacific (Pederson, 1983; Pederson et al, 1988; Herguera & Berger, 1991; Herguera, 1992;1994; Thunell et al., 1992; Berger et al., 1994). In the more complex monsooninfluenced, upwelling area of the Northern Indian Ocean, primary productivity was higher during interglacial than glacial periods of isotopic stages 11 until 2 but the signal was reversed during isotopic stages 13 and 12 (Steens et al., 1992). With these geographic differences in glacial/interglacial primary productivity estimates, it is important to have a global ocean coverage to come up with a primary productivity signal to be used in global carbon models. Organic carbon Although organic carbon is extensively used as a primary productivity proxy in upwelling areas, it is unreliable in the studied open-ocean area because of the very low organic carbon content of the sediments and the contamination by terrigenous organic carbon. The very low organic carbon, in addition to the high carbonate content of the sediments resulted to the poor reproducibility of the data (Fig. 11) even though the method adapted for this study (Nieuwenhuize et al, in press), eliminates reweighing and therefore minimises losses of organic compounds. Aside from being low, the total organic carbon is contaminated by terrestrial organic material. The high weight percent total organic carbon in stages 4-2 coincides with high abundances of land-derived biomarkers consisting of n-alkanes (C 27, C 2 9 and C 31), n-alkanoic acids (C 24, C 2 6 and C 28) and n-alkanols (C 24, C 2 6 and C 28) analysed in the same core (Madureira et al., in press). There is also a maximum in terrestrial biomarker abundances in h3 and h6, corresponding to high organic carbon content. This reflects increased input of terrigenous organic matter partly by icebergs which are known to have brought in material from Canada during Heinrich events hi, h2,

30 52 van Kreveld. Planktic Ibiarninifera as primary productivity and water mass indicators,scriptageol, ) h4, and h5 (Grousset et al., 1993). During h3, the land-derived organic matter may have been transported by icebergs originating from Europe (Grousset et al., 1993). Aside from icebergs, the stronger winds during glacial periods (Fuhrer et al., 1993, GRIP Members, 1993) may have also brought in land-derived organic matter to the area (Madureira et al., in press). The high weight percent organic carbon in stages 4-2 may also reflect better preservation rather than productivity variation. Organic carbon is severely affected by heterotrophic activity such that only a very small portion of the primary production in the euphotic zone is eventually preserved in the sediments. The lowered concentration of bottom water oxygen during the last glacial period (Boyle & Keigwin, 1982; Curry & Lohmann, 1983), likewise lowered heterotrophic activity on the seafloor and possibly enhanced organic carbon preservation (Emerson, 1985). Better preservation by rapid burial may also account for the higher glacial organic carbon content (Müller & Suess, 1979), as the sedimentation rate for this core is about 2 to 3 times higher in isotopic stages 4-2 than the other stages. Organic carbon preservation is highly complex and has the potential for contamination by terrestrial organic matter making interpretations difficult, particularly in this open-ocean site. Water masses Since water masses have characteristic assemblages of biota, it is possible to recognise water mass variations based on changes in these organisms (Boltovskoy, 1959, 1962). Bé & Tolderlund (1971) and Bradshaw (1959), who studied the distribution of planktic Foraminifera in more than 700 plankton tows in the Atlantic and Pacific oceans showed that assemblages of planktic Foraminifera indeed characterise ocean water masses as defined by temperature and salinity. Ottens (1991) used cluster analyses and diversity indices to relate foraminiferal assemblages to hydrographically defined water masses. The distribution and abundance of planktic Foraminifera in sediment core top samples by analogy with modern distributions, also reflect water masses (Imbrie & Kipp, 1971; Kipp, 1976), therefore, planktic Foraminifera were used to trace water mass movements in the Quaternary (Ruddiman & Mclntyre, 1976; Mclntyre et al., 1972). Planktic Foraminifera species are used to trace water mass changes during the past 208 ka by grouping the species using cluster and correspondence analyses, then comparing these groups to living plankton distributions. Although there are some detailed dissimilarities, the observed groups are similar to the distribution of living plankton reported by Bé & Tolderlund (1971), Bé (1977) and Ottens (1991). Clusters I to III are interpreted to reflect polar, subpolar, and North Atlantic Current (NAC) water masses, respectively. Species of clusters IV and V, which constitute <4% of the total foraminiferal assemblage, may reflect transitional and subtropical water masses. Neogloboquadrina pachyderma (s), which is interpreted to indicate polar water masses, dominates present day polar waters of the Arctic and the Antarctic oceans. This species can live in sea ice in both the northern (Carsola, 1953) and the southern (Spindler & Dieckmann, 1986; Dieckmann et al., 1991) hemispheres, and can tolerate

31 van Kreveld. Planktic Foraminiiera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 53 temperatures of -1.4 C though it favours temperatures between 0-9 C (Bé & Tolderlund, 1971). It dominates, reaching between % of the foraminiferal assemblage, in net tow samples from the Nansen Basin (Carstens & Wefer, 1992) and sediment samples from Greenland Sea (Bauch, 1993), Fram Strait and Central Arctic Ocean (Ericson, 1959). Turborotalita quinqueloba, which is designated as a subpolar species, has the highest absolute and relative frequencies in present day subpolar waters (Bé & Hamlin, 1967; Bé & Tolderlund, 1971; Ottens, 1991). It also dominates sediment trap samples (Reynolds & Thunell, 1985) and surface sediments in subarctic regions (Kipp, 1976). This species prefers surface temperatures between 12 and 15 C (Bé & Hamlin, 1967). Cluster III species are interpreted to indicate the NAC water mass (similar to 'transitional' of Bé & Hamlin, 1967), which can be further subdivided into two subclusters; (1) G. bulloides, N. incompta and G. scitula and (2) G. glutinata and G. inflata. Present day NAC water is a mixture of cold and warm waters which is reflected in the fauna. Aside from being prolific in NAC, cool water species G. bulloides, N. incompta and G. scitula also abound in subpolar waters while G. glutinata is quite common in subtropical waters (Bé & Hamlin, 1967; Bé & Tolderlund, 1971). Globorotalia inflata is also interpreted as a NAC indicator, agreeing with its living distribution, being abundant in the boundary region between Labrador Current and the Gulf Stream-North Atlantic Current, and common in northern east Atlantic (Bé & Hamlin, 1967; Bé & Tolderlund, 1971). Species of clusters IV and V are found in present day transitional to subtropical water masses. Discrepancies between my Foraminifera-based water mass designations, and the living and sea-bed plankton distributions can be explained by differences in sieve size and depth habitats sampled. I analysed the >125 urn fraction while Bé & Tolderlund (1971), and Bé & Hamlin (1967), examined the >200 urn fraction, therefore underestimating the contribution of smaller Foraminifera such as G. scitula and T. quinqueloba while overestimating larger ones. Kipp (1976) analysed the >150 urn while Barash (1971) counted the >100 urn fraction of sediment samples. Since Foraminifera live at different depths in the water column (Vincent & Berger, 1981), species distributions based on surface plankton only (e.g. Bé & Tolderlund, 1971; Ottens, 1991) underrepresents deep-living species such as G. scitula which is well represented in sediment samples (Kipp, 1976). Moreover, data on living plankton represent only several hours to days while sediment data record an average signal over at least hundreds or even thousands of years incorporating both seasonal and interannual variations. This is illustrated in the transitional boundaries between the designated water masses (Fig. 16A-B). Although plankton tow and sediment data are not directly comparable, the broad patterns are very similar. Therefore, Foraminifera in the sediments can still be used to reconstruct past water mass changes. Diversity indices are also used to characterise present day water masses (Ottens, 1991). The Shannon diversity index for polar samples is low to very low, for subpolar samples moderate, and high for NAC to subtropical samples (Fig. 18). The overall pattern is comparable to box-core top data which show a decrease in planktic foraminiferal diversity from low to high latitudes in the Atlantic Ocean, generally agreeing

32 54 van Kreveld. Planktic Foraminifera as primary oductivity and water mass indicators, Scripta GeoL, ) with modern sea-surface temperature and circulation (Balsam et al., 1980). All three diversity indices, simple, Shannon, and equitability, are distinctly low during Heinrich events coinciding with the dominance of polar waters. The low equitability values are due to the almost monospecific planktic foraminiferal assemblage of N. pachyderma (s). The foraminiferal assemblages characteristic of particular water masses have shifted through time in relation to the location of the piston core (Figs. 14A-B). The area is interpreted to be dominantly bathed by NAC with minor incursions of transitional and subtropical waters in interglacial periods and by polar waters during Heinrich events. During glacial periods, the area was covered by polar waters succeeded by subpolar waters and vice-versa. The water mass reconstructions agree well with those of Mclntyre et al. (1972), Ruddiman & Mclntyre (1976), CLIMAP (1981), and Crowley (1981). Productivity and water masses The reconstructions generally show moderate to high primary productivity during interglacial periods when NAC water predominantly covered the area (compare Figs. 7 and 14A-B). At present, this site is mostly bathed by the NAC, the westerly wind-driven Gulf Stream extension. When the westerlies are strong during the cold season, they induce vertical mixing, bringing nutrients to surface waters. Enough nutrients and optimal light support algal spring blooms (Holligan et al., 1983; Brown & Yoder, 1994a, b) and high concentrations of planktic Foraminifera (Bé & Hamlin, 1967) currently observed in this area. Modern polar waters in the Arctic as well as Antarctic generally have low productivities (Piatt & Subba Rao, 1975; Subba Rao & Piatt, 1984) which are comparable to those during Heinrich events. Icebergs probably blocked sunlight and caused a strong salinity stratification in the surface water suppressing admixture of nutrients and plankton productivity. The moderate productivity during glacial periods may be due to localised blooms when the ice thawed, similar to present day ice-edge environments (Hebbeln &Wefer,1991). Conclusions Piston core T88-9P was recovered from the Northeast Atlantic at 48 23'2"N, "W and provides a continuous pelagic record unaffected either by carbonate, dissolution or by turbidity currents because the core site lies well above the lysocline at 3193 m water depth and on the east flank of the mid-atlantic ridge, far from continental slopes where turbidity currents are common. The age model for this core reveals a 208 ka stratigraphie record back to the beginning of isotopic stage 7, based on dates assigned to isotopic stage boundaries and events, calendar-calibrated radiocarbon ages, and tephrochonology. The average total planktic Foraminifera are generally higher in interglacial ( specimens/g and specimens/emzka) than in glacial sediments ( specimens/g and specimens/cmzka, maximum estimate), and low (9200 spec-

33 van Kreveld. Planktic Foraminifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 55 Fig. 18. Scatter plot of % Neogloboquadrina pachyderma (s) against Shannon diversity index, with species clusters (I-V). NAC stands for North Atlantic Current. imens/g and specimens/cm ka, minimum estimate) in Heinrich layers. This 2 pattern is interpreted to reflect variations in foraminiferal productivity as a consequence of changes in surface water primary productivity. In general, primary productivity variations during the past 208 ka coincide with water mass changes. As in the modern oceans, primary productivity is generally higher in NAC and subpolar than in polar waters. The high average primary productivity during interglacials coincides with times when NAC water mostly covered the area as shown by the dominance of G. bulloides, N. incompta, G. scitula, G. glutinata, and G. inflata group in the sediments, while the photic zone had a low fertility during Heinrich events, when polar waters penetrated the area as reflected by the low diversity indices and the predominance of N. pachyderma (s). Productivity was moderate during glacial periods when polar, subpolar and NAC waters, probably in seasonal alternation, covered the area as reflected by the abundance of subpolar species T. quinqueloba as well as polar and NAC indicator species in the sediments. The very low weight percent total organic carbon and the contamination by landderived organic matter limit the use of organic carbon as a primary productivity proxy in the study area. In contrast, planktic Foraminifera are useful tracers for past

34 56 van Kreveld. Planktic Foraminifera as primary productivity and water mass indicators,scriptageol, ) primary productivity and water mass changes in the carbonate-dominated, openocean, mid-latitude Northeast Atlantic site. Acknowledgements I am grateful to L. Labeyrie and E. Cortijo for digitising the core photograph. J.E. van Hinte and G.M. Ganssen initiated this research. They are also thanked for critically reviewing several versions of this manuscript along with M. Knappertsbusch and J.J. Ottens. I thank S. Kars for taking the Scanning Electron Micrographs and A.J. Nederbragt for providing the cluster and correspondence analyses programs and for her useful suggestions. Thanks are also due to M. Konert, R. van Elsas and R Willekes for analysing the organic carbon. This study is a part of the GEM program (Global Emiliania Modelling Initiative) coordinated by P. Westbroek and is financed by the 'Nederlandse Onderzoek Project'. The APNAP II expedition which collected the studied piston core was sponsored by the 'Stichting Onderzoek der Zee'. This is publication number of the Netherlands Research School of Sedimentary Geology, Center for Marine Earth Sciences, Free University, Amsterdam. References Abrantes, F., K. Winn & M. Sarnthein, Late Quaternary palaeoproductivity variations in the NE and equatorial Atlantic: diatom and C o r g evidence. In: R. Zahn, T.H. Pederson, M.A. Kaminski, and L. Labeyrie (eds.). Carbon cycling in the glacial ocean: constraints on the ocean's role in global change. NATO ASI Series, 117: Anderson, O.R., M. Spindler, A.W.H. Bé & C. Hemleben, Trophic activity of planktonic Foraminifera. J. Mar. Biol. Assoc. U.K., 59: Andrews, J.T. & K. Tedesco, Detrital carbonate-rich sediments, northwestern Labrador Sea: implications for ice-sheet dynamics and iceberg rafting (Heinrich) events in the North Atlantic. Geology, 20: Balsam, W.L., K.W. Flessa, N.G. Kipp & L.G. Dubois, Planktonic foraminiferal diversity in the interglacial and glacial North Atlantic: a test of diversity gradients as a paleoceanographic technique. Geology, 8: Barash, M.S., The vertical and horizontal distribution of planktonic Foraminifera in Quaternary sediments of the Atlantic ocean. In: B.M. Funnel & W.R. Riedel (eds.). The Micropaleontology of the oceans. Cambridge Univ. Press, London: Bard, E., Correction of accelerator mass spectrometry C ages measured in planktonic Foraminifera: paleoceanographic implications. Paleoceanogr., 3,6: Bard, E., M. Arnold, R. Fairbanks & B. Hamelin, Th-2^U a nd "C ages obtained by mass spectrometry on corals. Radiocarbon, 35: Bard, E., B. Hamelin, R. Fairbanks & A. Zindler, Calibration of the C timescale over the past ,000 years using mass spectrometric U-Th ages from Barbados corals. Nature, 345: Barnola, J.M., D. Raynaud, Y.S. Korotkevich & C Lorius, Vostok ice core provides 160,000-year record of atmospheric C02. Nature, 329: Bauch, H., Planktische Foraminiferen im europäischen Nordmeer - ihre Bedeutung für die paläo-ozeanographische Interpretation während der letzten Jahre. Sonderforschungsbereich, 313, Univ. Kiel, Germany, 40: Bé, A.W.H., Ecology of Recent planktonic Foraminifera 1. Areal distribution in the western North Atlantic. Micropaleontology, 5: Bé, A.W.H., An ecological, Zoogeographie, and taxonomie review of recent planktonic Foraminifera. In: A.T.S. Ramsay (ed.). Oceanic Micropaleontology, 1. Academic Press, London:

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41 van Kreveld. Planktic R)rarninifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 63 Appendix 1 Annotated alphabetical species list Plates 1-12 represent Scanning Electron Micrographs of foraminifera specimens from the Northeast Atlantic piston core T88-9P. RGM stands for the National Museum of Natural History, Palaeontology Department, formerly Rijksmuseum van Geologie en Mineralogie, Leiden. I follow the taxonomy of Bé & Tolderlund (1971) with some modifications by Saito et al. (1981). Globigerina bulloides d'orbigny, 1826 PI. 1, figs In the North Atlantic, G. bulloides is a dominant species in NAC waters (Ottens, 1991,1992) and abounds in subpolar waters as well, from New York to Portugal and from the Labrador to the Norwegian Sea (Bé & Hamlin, 1967; Bé & Tolderlund, 1971). It has a similar distribution in surface sediment samples (Kipp, 1976). G. bulloides is also a characteristic species in upwelling regions (Prell & Curry, 1981; Prell, 1984a,b). It occurs over a surface temperature range of 0 C to 27 C (Bé & Tolderlund, 1971). G. bulloides has high (up to 29%), moderate and low accumulation rates, relative and absolute frequencies in interglacial, glacial and Heinrich layers, respectively. The species is about twice as frequent in interglacial as in glacial sediments. Globigerina rubescens Hofker, 1956 Pl. 1, figs Most specimens were white although some had a pink/orange pigmentation. This species is distinguished from G. tenellus by its pigmentation and by the absence of a secondary aperture(s). This species may comprise up to 2.3% in interglacial, is seldom present in glacial and absent in Heinrich layers. In the 208 ka record, the maximum relative and absolute frequencies and accumulation rate of Globigerina rubescens are noted immediately after Termination II, just like those of the other warm-water species such as Globigerinoides tenellus and Orbulina universa. Globigerinella aequilateralis (Brady, 1879) PI. 2, figs G. aequilateralis is only present in low frequencies not exceeding 2.3% in interglacial, seldom present in glacial and generally absent in Heinrich sediments. There are two forms, the loosely-coiled being more common than the involute and tightlycoiled tests. Peak absolute frequency of 490 specimens/g is noted in isotopic stage 7.

42 64 van Kreveld. Planktic R)raminifera as primary productivity and water mass indicators,scriptageol, ) Globigerinita glutinata (Egger, 1893) PL 3, figs The supplementary features like bullae and secondary apertures give G. glutinata many morphologic variations even within a single sample. The relative frequency of this species is high (reaching 20%) in interglacial, moderate in glacial and very low in Heinrich layers. The mean absolute frequency is 4800, 1000 and 400 specimens/g, respectively. The interglacial accumulation rate is generally higher than the glacial. Globigerinita uvula (Ehrenberg, 1861) This very small microperforate species with a high trochospiral and translucent, smooth test is identified only in sample 50 at about 12.5 ka. The accumulation rate, relative and absolute frequencies are 3090 specimens/cm ka, <1%, and 780 specimens/g, 2 respectively. Globigerinoides ruber (d'orbigny, 1839) PL 4, figs There are three G. ruber forms: (1) a small, compact, low-spired, thick- walled form; (2) a bigger, moderately high-spired and lobular variant; and (3) a high-spired and thin-walled form. G. ruber also exhibits two colour varieties, pink and white. The pink form is generally larger than the white one and usually occurs in warmer waters in the Atlantic, whereas the white form prefers lower temperatures (Hemleben et al., 1989). Most specimens are white except for a few pink ones in isotopic substage 5.5. This species is absent from isotopic stages 2 and 3, and does not exceed 3% of the total planktic foraminiferal assemblage in the other isotopic stages. Both the average absolute frequency and the accumulation rate are higher in interglacial than in glacial periods. Globigerinoides tenellus Parker, 1958 PL 5, figs G. tenellus was distinguished from G. rubescens by the presence of dorsal supplementary aperture(s) and from G. ruber by its more lobulate periphery. This species may comprise up to 3.7% of the planktic foraminiferal assemblage in interglacial sediments. It is absent from Heinrich and most glacial sediments. Globorotalia hirsuta (d'orbigny, 1839) PL 6, figs G. hirsuta is a cool subtropical species (Ruddiman & Mclntyre, 1976). Like Globorotalia truncatulinoides it reproduces at the surface and then sinks to deeper waters where it prefers to live (Hemleben et al., 1989).

43 van Kreveld. Planktic Ibrarninifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 65 This species can be big with a relatively high spire, pustulate surface and a weakly lobulate periphery in interglacial sediments while glacial specimens are small with a relatively lower spire. G. hirsuta is absent from Heinrich layers and isotopic stages 2 and 4, and occurs in relatively low amounts (not exceeding 3%) in the other stages. Its average absolute frequency and accumulation rate are about four times higher in interglacial than in glacial sediments. Globorotalia inflata (d'orbigny, 1839) PI. 7, figs G. inflata abounds in the Gulf Stream and the North Atlantic Current (Bé & Hamlin, 1967; Bé & Tolderlund, 1971) but inhabits subpolar waters as well. It is a deepdweller preferring depths between m (Bé, 1977). Most specimens in the core have a smooth and shiny outer layer with a convex spiral side and rounded periphery. However, some specimens have a granular texture and subangular periphery. This species has high, moderate and low frequencies and accumulation rates in interglacial, glacial and in Heinrich layers, respectively. G. inflata may comprise up to 16.0 % of the foraminiferal assemblage with a maximum absolute frequency of 4170 specimens/g in interglacial sediments. Globorotalia scitula (Brady, 1882) PI. 2, figs G. scitula is found in subpolar waters in rather high frequencies (Rea et al., 1991; Reynolds & Thunell, 1985) and in subtropical waters (Bé & Hamlin, 1967). The relative frequency of G. scitula is higher (reaching 8.5%) in interglacial than in glacial sediments. Both absolute frequency and accumulation rate are also generally higher in the former than in the latter. It occurs in very low frequencies or may be absent in Heinrich layers. Globorotalia truncatulinoides (d'orbigny, 1839) PI. 6, figs G. truncatulinoides is a subtropical species (Bé & Tolderlund, 1971). It is common in the Sargasso Sea where it reaches a maximum abundance during winter (Bé, 1959; Bé & Tolderlund, 1971). This species occurs in low accumulation rates, absolute and relative frequencies (not exceeding 3.7%) in interglacial layers. Its peak absolute frequency is at 126 ka, immediately after Termination II. It is absent from isotopic stage 2 and in most of stages 6,4 and 3. Neogloboquadrina incompta (Cifelli, 1961) PI. 8, figs N. incompta has been regarded as a dextral ecophenotype of N. pachyderma or a

44 66 van Kreveld. Planktic Ibraminifera as primary productivity and water mass indicators,scriptageol, ) dextral-coiling subspecies thereof (Parker, 1962; Reynolds & Thunell, 1986). I followed the classification of Ciffeli (1961, 1973) and distinguished N. incompta by its dextral coiling, more pustulate surface and more lobulate form compared to N. pachyderma. The relative frequency of N. incompta is high (reaching 33% in isotopic stage 5), moderate (up to 23 %) and very low in interglacial, glacial and Heinrich layers, respectively. Both absolute frequency and accumulation rate are higher in interglacial than in glacial sediments. Neogloboquadrina pachyderma (Ehrenberg, 1861) Pl. 9, figs This species inhabits polar as well as subpolar waters (Bé & Hamlin, 1967), can live within sea ice and tolerates salinities of up to 82%o (Hemleben et al., 1989). N. pachyderma was initially separated into two groups A and B, with group A consisting of crystalline forms which results from a secondary calcification process over a primary reticulate surface texture, and group B having reticulate textures (Srinivasan & Kennett, 1974). Reynolds & Thunell (1986) adapted the same classification and added that group A is predominantly sinistral while group B is predominantly dextral. For this study, Reynolds & Thunell's (1986) group A was identified as N. pachyderma. Bé & Hamlin (1967) demonstrated that the distribution boundary between dextral and sinistral forms (according to my classification, N. incompta and N. pachyderma, respectively) is associated with the 7.2 C surface isotherm. Aside from temperature changes, coiling patterns could also reflect availability of nutrients with dextral forms abundant when nutrient concentration is low and sinistral forms being abundant when nutrient concentration is high (Reynolds & Thunell, 1986). However, Brummer & Kroon (1988) showed that changes in coiling patterns reflect the dynamics of central water masses and polar fronts rather than the displacements of particular isotherms. There are two major N. pachyderma forms in the core. One has a small compact test with a rounded outline, and has thick walls which tend to obscure the sutures and the aperture. The other has a large lobulate test and 4 to 5 discrete chambers separated by sutures, with the last chamber reduced or bullate, sometimes partially covering the aperture. Both forms are abundant in Heinrich and glacial layers while the former is more common in interglacial sediments. The relative frequency of N. pachyderma is low (not exceeding 20%), moderate and very high (up to 95%) in interglacial, glacial and Heinrich layers, respectively. Despite its dominance in Heinrich layers, its absolute frequency is generally lower compared to glacial sediments. Its maximum absolute frequency and accumulation rate is in isotopic stage 6. Orbulina universa d'orbigny, 1830 PI. 5, figs The size of this species is correlated with the availability of food supply, the more

45 van Kreveld. Planktic Foraminifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 67 food, the larger the spherical shell and overfeeding results in a double sphere, a socalled 'Biorbulina' (Hemleben et al., 1989). There are some large tests in interglacial sediments. This species may comprise up to 2% of the total planktic foraminiferal assemblage in interglacial sediments. It is absent from glacial sediments except in isotopic stage 6 where it is occasionally present. Turborotalita humilis (Brady, 1884) PL 5, figs T. humilis is found only in sample number 715 cm, at 192 ka. It has a very low frequency of 0.2 %, corresponding to 360 specimens/cm 2 ka. This species differs from T. quinqueloba by its smaller size and more numerous chambers. Turborotalita quinqueloba (Natland, 1938) PL 10, figs. 1-14; PL 11, figs. 1-18; PL 12, figs This typically subpolar species (Bé & Hamlin, 1967; Bé & Tolderlund, 1971) shows considerable variation in the core. Numerous specimens have a bulla-like elongation of the final chamber which can partially or completely cover the aperture, others have an aperture without a lip, while some have a thickened rim-like basal hp. The relative frequency of T. quinqueloba is high in interglacial (reaching 31.9%) and in glacial (reaching 38.8%) sediments while it is low in Heinrich layers. The average absolute frequency and accumulation rate of this species are higher in interglacial than in glacial sediments.

46 68 van Kreveld. Planktic Foraminifera as primary productivity and water mass indicators,scriptageol., ) Plate 1 The 100 urn bar scale below fig. 2 applies to figs. 1-6; the scale on the lower left corner is for figs. 8-12; and the 10 urn bar is for figs. 7 and 13. Figs Globigerina bulloides d'orbigny, Umbilical views. Specimens of figs. 1, 5, and 6 are from sample 0, of figs. 2-4 and 7 are from sample Typical specimen with lobulated periphery and four spherical to subspherical chambers in the final whorl; RGM Common morphotype with wide apertural opening; RGM Thinner-shelled, smoother-surfaced specimen with chambers rapidly increasing in size; RGM Specimen with diminutive ultimate chamber in the final whorl; RGM Spiral view; RGM Side view; RGM Detail of surface wall structure showing large irregularly-spaced pores. Elevated areas are spine bases, disappearance of spine shafts reveal circular holes in the centre; RGM Figs Globigerina rubescens Hofker, 1956 from sample Spiral view; note absence of secondary aperture; RGM Side view; RGM Umbilical view of same specimen in fig. 8 showing wide highly-arched aperture. 11. Umbilical view of specimen with thick-wall, compact test and small aperture with distinct lip; RGM Spiral view; RGM Same specimen as in fig. 8 showing last chamber to have shallow depression leading to roughly equidistant pores. Pores are small and interpore areas are wide.

47 van Kreveld. Planktic Foraminifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 69

48 70 van Kreveld Planktic R)rarninifera as primary productivity and water mass indicators, ScriptaGeoL, ) Plate 2 The 100 urn scale bar applies to figs. 1-9 and figs , the 10 urn bar to figs. 7 and 12, and the 1 urn bar to fig. 13. Figs Globigerinella aequilateralis (Brady, 1879). Specimens of figs. 1-3 and figs. 5-7 are from samples 0 and 500, respectively. 1. Umbilical view of large, loosely-coiled specimen; note smooth surface of last chamber compared to rest of test; RGM Spiral view showing partial separation of last two chambers; RGM Asymmetricallycoiled specimen with crescent-shaped aperture; RGM Umbilical view of small, tightly-coiled specimen; RGM Spiral and side views of small specimens displaying asymmetrical coiling; RGM Enlargement of last chamber of specimen in fig. 4 shows spine bases as upraised circular to triangular mounds with round holes. Living specimens have triradiate spines. Figs Globorotalia scitula (Brady, 1882) specimens from sample Spiral view; note large pores with wide pore pits within and around the proloculus, decreasing in size in older chambers; RGM Side view showing irregularly-shaped lip; RGM Spiral view of specimen with slowly increasing chamber size giving a nearly circular peripheral outline; RGM Umbilical view of a specimen that is heavily pustulate near the apertural area; RGM Enlargement showing smooth finely perforate last chamber of specimen in fig Further enlargement of a single pore of fig. 12 illustrates that widening of pore size as the wall thickens with age, results to a well-layered conical pit.

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50 72 van Kreveld. Planktic Foraminifera as primary productivity and water mass indicators,scriptageol, ) Plate 3 This plate illustrates the extreme variability in Globigerinita glutinata (Egger, 1893) test morphology. The 100 um scale bar applies to figs Figs. 5, 11 and 15 are specimens from sample 0 while the rest are from sample 500. Figs Umbilical views. 1. Simple primary aperture with thin lip; RGM Specimen with inflated last chamber and flap covering aperture; RGM Circular bulla covers aperture; RGM Specimen with dumbbell-shaped bulla; RGM Large inflated bulla with four infralaminal apertures; RGM Complex bulla with apertures at sutures of older chambers; RGM Intricate bulla with multiple apertures covering considerable part of test; RGM Figs Spiral views. 8. Specimen without secondary aperture; RGM Specimen with one secondary aperture. I consider this form to be synonymous to Globigerinoides parkerae Bermudez; RGM Specimen with large secondary aperture almost divided into two parts; RGM Infralaminal aperture of specimen extends to spiral side; RGM Part of bulla and two infralaminal apertures reach spiral side RGM Figs Side views. 13. Slit-like opening on the side; RGM Specimen with secondary aperture; RGM Smooth and thin-walled specimen with irregularly defined bulla that has multiple infralaminal apertures; RGM Figs Detailed views. 16. Specimen in fig. 13 showing bulla with smaller pustules than the remainder of the test. 17. Bulla of specimen in fig. 7 displaying conical pustules and small, dense but irregularly-spaced pores on their bases. 18. The two infralaminal apertures of fig. 12 specimen open beneath the bulla.

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52 74 van Kreveld. Planktic Foraminifera as primary productivity and watermassindicators,scriptageol,113(1996) Plate 4 Test morphology and microstructure of Globigerinoides ruber (d'orbigny, 1839). The 100 urn bar scale applies to figs Figs. 10 and 14 specimens are from sample 500 and the rest are from sample 15. Figs Small, compact and thick-walled form. 1. Apertural view; RGM Spiral view showing one accessory aperture; RGM Side view; RGM Figs Larger, more lobulate form Umbilical views of specimens having circular apertures with and without a lip, respectively; RGM Specimens with one and two apertures on the spiral side, respectively; RGM Side view showing an accessory aperture; RGM Figs Large and high trochospiral form. Side views. 9. Specimen with small last chamber; RGM Thin-walled with inflated chambers rapidly increasing in size. Note same magnification as other G. ruber variants; RGM Fig Detailed views. 11. Penultimate chamber of fig. 1 specimen showing pores between mostly isolated and few interconnected spine bases with circular spine holes. 12. Last chamber of fig. 6 specimen illustrating big, inequidistant pores and spine bases. 13. Ultimate chamber of fig. 9 specimen shows large cone-shaped pores. 14. Last chamber of fig. 10 specimen showing rounded spine bases with circular holes.

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54 76 van Kreveld. Plankrk Foraminifera as primary productivity and water massindicators,scriptageol,113(1996) Plate 5 The 100 urn scale bar on the upper left applies to figs. 1-3 and 6-7 while the lower left one is for figs The 10 urn scale bar applies to figs. 4,5 and 11. Figs Globigerinoides tenellus Parker, 1958 specimen from sample Umbilical view of specimen with four spherical chambers in the final whorl and circular, highlyarched primary aperture with faint rim-like lip; RGM Spiral view of fig. 1, showing two accessory apertures, big and small. 3. The big accessory aperture is still visible from the side; RGM Detail of last chamber of fig. 1, illustrating coarsely perforated wall structure with large pores set in shallow depressions. Pore depressions are bounded by weakly developed polygonal ridges and spine bases are located at the junction between these ridges. Disappearance of the spine shafts show round spine holes. Figs Orbulina universa d'orbigny, 1830 specimens from sample Wall structure of fig. 6 specimen shows big and small pore openings of about 15 and 5 urn, respectively. The small pores of living specimens possess an inner organic lining and pore plates (Bé, et al, 1980) while the large pores lack these structures, and are therefore defined as apertures. 6. Big specimen with irregularly-spaced areal apertures; RGM Small specimen showing same feature; RGM Figs Turborotalita humilis (Brady, 1884) specimen from sample Umbilical view showing 6 1/2 chambers in the last whorl, with ultimate chamber extending basally to cover aperture; RGM Spiral view of fig. 8; note deeply-incised sutures. 10. Side view of fig Enlargement of part of ultimate and penultimate chambers showing intergrowth of angular, euhedral calcite crystals to conceal some pores; same specimen.

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56 78 van Kreveld. Planktic Foraminifera as primary productivity and watermassindicators,scriptageol,113(1996) Plate 6 The 10 urn bar scale applies to figs. 4 and 5 while the 100 urn bar to the rest. Figs Globorotalia hirsuta (d'orbigny, 1839) from sample Umbilical view displaying intergrowth of nodelike pustules forming botryoidal surface near apertural area. Most pores in earlier chambers are obliterated by abundant interlocking pustules except in ultimate chamber; RGM Spiral view showing angular, platelike structures thickening prolocular area instead of knoblike pustules seen on spiral side; RGM Side view; RGM Close-up of last chamber of fig. 1 specimen showing only few isolated pustules. Note uneven surface texture due to partial calcite layer cover. Figs Globorotalia truncatulinoides (d'orbigny, 1839) from sample Enlargement of last chamber displaying medium-sized, dense pores and nodelike pustules in flat surface; RGM Umbilical view. As in G. hirsuta, test is very thick near aperture and thins in direction of growth, with little thickening in last chamber; RGM Spiral view; note big dense pustules in prolocular area and at keel junction; RGM Side view showing triangular peripheral outline and aperture with lip; RGM

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58 80 van Kreveld. Plankticforaminiferaas primary productivity and water massindicators,scriptageol,113(1996) Plate 7 Test morphology and wall microstructure of Globorotalia inflata (d'orbigny, 1839). The 10 urn bar scale applies to figs. 10 and 11 while the 100 urn bar to the rest. Figs. 1-3 are specimens from sample 0 while figs are from sample 450. Figs Large thick-walled form. 1. Umbilical view of typical specimen which is coarsely granular in umbilicus and smooth in last chamber; RGM Spiral side has smooth wall structure; RGM Side view shows flatter spiral side of specimen compared to more convex ones of specimens in figs. 6 and 9; RGM Figs Large, thick-walled form with diminutive last chamber. 4. Umbilical view; note smooth diminutive and penultimate last chamber; RGM Spiral view showing almost rounded peripheral outline; RGM Side view displays large pustules around umbilicus; RGM Figs Small, thin-walled forms. 7. Umbilical view; RGM Spiral view illustrates rougher wall texture compared to specimens in figs. 2 and 5; RGM Side view shows more angular outline due to protrusion of last chamber; RGM Fig Enlarged views. 10. Penultimate chamber of specimen in fig. 5 has smooth, hummocky wall structure with constricted or obliterated pores. Most pores are on top of hummocks but few are located in depressions in between. 11. Last chamber of specimen in fig. 8 has porous surface with discrete nodelike pustules; note bigger irregularly-spaced pores compared to specimen with veneer of calcite crust (fig. 10).

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60 82 van Kreveld. Planktic Foraminifera as primary productivity and water massindicators,scriptageol.,113(1996) Plate 8 Morphological variants of Neogloboquadrina incompta (Cifelli, 1961) from samples 0 (fig. 2), 702 (figs. 1, 6 and 10) and 450 (figs. 3-5, 7-9 and 11-12). The 100 urn scale bar applies to all figures except for fig. 5. Figs Umbilical views. 1. Lobulate specimen with 5 chambers in last whorl separated by distinct sutures. Chambers rapidly increase in size as added with last chamber partially covering aperture; RGM Less lobulate specimen with bean-shaped last chamber totally covering aperture; RGM Specimen has deep umbilicus and almost circular peripheral outline with the chambers slowly increasing in size as added (compare to fig. 1); RGM Specimen with four chambers in last whorl; ultimate chamber has flap with lip partially covering aperture; RGM Fig. 5. Enlarged view of penultimate chamber of specimen in fig. 7 showing large pores with calcite ridges surrounding individual pore pits. The sharp, uneven crystals of these ridges give an overall rough texture (compare with wall structure of N. pachyderma in PI. 9, figs ). Figs Spiral views. 6. Note the thickened prolocular area; RGM Specimen with quadrate peripheral outline and smaller ultimate than penultimate chamber; RGM Note large last chamber; RGM Figs Side views. 9. Note aperture with thick lip; RGM Specimen with wide apertural opening only partially covered by a lip; RGM Aperture without a lip; RGM Last chamber covers most of aperture; RGM

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62 84 van Kreveld. Planktic Foraminifera as primary productivity and watermassindicators,scriptageol,113(1996) Plate 9 Neogloboquadrina pachyderma (Ehrenberg, 1861) specimens from samples 500 (figs. 1-3), 70 (figs. 4-6 and 13) 100 (figs and 14-15). The 100 urn scale bar applies to figs while the 10 urn bar to figs Figs Small, very thick-walled, typical interglacial form. 1. Specimen with compactly arranged chambers. This is classified as N. pachyderma, group A by Reynolds & Thunell (1986); RGM Spiral view shows rounded test outline; RGM Side view. Note aperture is covered by last chamber; RGM Figs Large and very thick-walled specimens from a Heinrich layer. 4. Umbilical view. Although this form is similar to fig. 1, Heinrich layer and glacial specimens can attain big sizes. Note wide umbilicus surrounded by distinct calcite rhombs; RGM Spiral view showing sutures obliterated by heavy calcification; RGM Part of aperture still visible on side view; RGM Figs Quadrate morphotype typical of glacial sediments. 7. Umbilical view showing wide aperture covered by unornamented thin lip. A less encrusted specimen with similar form is classified by Reynolds & Thunell (1986) as N. pachyderma, group B; RGM Spiral view. Note sutures obliterated by calcite crust; RGM Side view showing wide apertural opening; RGM Figs Ovate morphotype common in glacial sediments. 10. Umbilical view of 4 1/2-chambered specimen with diminutive last chamber covering part of aperture; RGM Spiral view showing faint sutures; RGM Side view; note small last chamber; RGM Figs Enlarged views. 13. Last chamber of fig. 4 specimen showing calcite crust to obliterate some pores. 14. Wall structure and topography of penultimate chamber of fig. 8 specimen illustrating rugged ridges surrounding pores. 15. Penultimate chamber of fig. 11 specimen showing smoothened surface texture with most pores closed-off.

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64 86 van Kreveld. Planktk: Foraminifera as primary productivity and water massindicators,scriptageol,113(1996) Plate 10 Extreme variability in Turborotalita quinqueloba (Natland, 1938) morphotypes due to differences in test thickening, shape of the chambers and aperture. All specimens are from interglacial sample 450 and are dextrally-coiled, except for those shown in figs. 5 and 13. The 100 urn bar scale applies to all figures. Figs Umbilical views. 1. Typical specimen with five chambers in the last whorl and an umbilical aperture bordered by small lip; RGM More lobulate form with big aperture and no lip. This specimen has spine bases and holes distinguishing it from small specimens of N. incompta; RGM Umbilical aperture with small unornamented lip; RGM Flap partially covers aperture. Note less thickened, deeply-incised apertural area; RGM Slightly compressed specimen; RGM Flap-like extension of last chamber is partially fused to an earlier chamber in apertural area; RGM Large lobular variant; RGM Terminal chamber has two flap-like extensions; RGM Tear drop-shaped last chamber with irregular flap; RGM Tear drop-shaped last chamber with projecting lip covering aperture; RGM Six chambers in last whorl with diminutive ultimate chamber; RGM Aberrant terminal chamber is smaller than penultimate chamber; RGM Large specimen with bullate last chamber totally covering the aperture and parts of earlier chambers; RGM Huge and rare T. quinqueloba variant with exceptionally big last chamber and thick crescentshaped ornamented lip; RGM

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66 88 van Kreveld. Planktie Foraminifera as primary productivity and watermassindicators,scriptageol,113(1996) Plate 11 Turborotalita quinqueloba (Natland, 1938) morphotypes from glacial and Heinrich layers except for figs. 13 and 14 which are specimens from interglacial sediments (sample 450). Figs. 1-4 and 15; figs. 5-8,11, 16 and 17; and figs. 9-10, 12 and 18 are specimens from samples 100, 70 and 702, respectively. Specimens shown in figs. 1-6, and 17 are dextrally-coiled while the others are sinistrally-coiled. The 100 urn bar scale applies to all figures. Figs Umbilical views. 1. Note triangular lip partially covering aperture; RGM Terminal chamber with flap-like extension; RGM Coalescence of some spine bases to form ridges gives this specimen a rough surface wall texture; RGM Large and lobular with intricate bulla; RGM Small compressed specimen with semi-circular aperture and thin lip; RGM Small with faint sutures due to calcite encrustation on surface; RGM Note wide aperture and irregularly-shaped unornamented lip; RGM Umbilical-interiomarginal aperture partially covered by an unornamented lip; RGM Small, lobulate with spherical chambers and circular umbilical aperture; RGM Note deeply-incised sutures particularly near the aperture; RGM Small specimen with big last chamber; RGM Note smooth surface compared to other specimens; RGM Figs Spiral views. 13. Large specimen with spherical chambers rapidly increasing in size as added; RGM Ultimate is smaller than penultimate chamber (compare with fig. 13); RGM Note distinctly smoother last chamber compared to other chambers; RGM Small specimen with smoother wall structure than those of figs ; RGM Small, compact specimen with large pores; RGM Note thickened prolocular area; RGM

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68 90 van Kreveld. Planktic Foraminifera as primary productivity and watermassindicators,scriptageol,113(1996) Plate 12 Turborotalita quinqueloba (Natland, 1938). Figs. 1-2, 3, 4 and 5 are specimens from sample 450,100, 702 and 70, respectively. The 100 urn scale bar applies to figs. 1-5 while the 10 urn bar to figs Figs Side views. 1. Large specimen with lip projecting away from aperture; RGM Part of lip is attached to an earlier chamber; RGM Lip projects inward partially closing aperture; RGM More oblong and less inflated specimen than those in figs. 1-3; RGM Small, compact with smooth texture; RGM Figs Enlarged views. 6. Penultimate chamber of specimen in PI. 10, fig. 6 showing medium-sized, irregularly-spaced pores and spine bases. 7. Terminal chamber of specimen in PI. 11, fig. 2; note pore doublets (a). 8. Less dense pores and spine bases in last chamber of specimen in PI. 11, fig. 4; note uneven coating of calcite. 9. Penultimate chamber of specimen in PI. 11, fig. 13 showing dense, medium- sized pores and rounded to subrounded spine bases with circular holes. 10. Ultimate and penultimate chambers of specimen in PL 11, fig. 5 shows calcite encrustation covered most spine holes and pores. Uncovered pores of this small specimen are big. 11. This small specimen (same as in PL 11, fig. 17; detail of penultimate chamber) has bigger and less dense pores than larger specimens of figs. 6 and 9.

69 van Kreveld. Planktk: Foraminifera as primary productivity and water mass indicators, Scripta GeoL, 113 (1996) 91